Molecular phylogeny of the diatom genera Amphora and Halamphora (Bacillariophyta) with a focus on morphological and ecological evolution

The taxonomic history of the diatom genus Amphora is one of a broad early morphological concept resulting in the inclusion of a diversity of taxa, followed by an extended period of revision and refinement. The introduction of molecular systematics has increased the pace of revision and has largely resolved the relationships between the major lineages, indicating homoplasy in the evolution of amphoroid symmetry. Within the two largest monophyletic lineages, the genus Halamphora and the now taxonomically refined genus Amphora, the intrageneric morphological and ecological relationships have yet to be explored within a phylogenetic framework. Critical among this is whether the range of morphological features exhibited within these diverse genera are reflective of evolutionary groupings or, as with many previously studied amphoroid features, are nonhomologous when examined phylogenetically. Presented here is a four‐marker molecular phylogeny that includes 31 taxa from the genus Amphora and 77 taxa from the genus Halamphora collected from fresh, brackish, and salt waters from coastal and inland habitats of the United States and Japan. These phylogenies illustrate complex patterns in the evolution of frustule morphology and ecology within the genera and the implications of this on the taxonomy, classification, and organization of the genera are discussed.     

Ptolemeba n. gen., a Novel Genus of Hartmannellid Amoebae (Tubulinea,Amoebozoa); with an Emphasis on the Taxonomy of Saccamoeba

Hartmannellid amoebae are an unnatural assemblage of amoeboid organisms that are morphologically difficult to discern from one another. In molecular phylogenetic trees of the nuclear‐encoded small subunit rDNA, they occupy at least five lineages within Tubulinea, a well‐supported clade in Amoebozoa. The polyphyletic nature of the hartmannellids has led to many taxonomic problems, in particular paraphyletic genera. Recent taxonomic revisions have alleviated some of the problems. However, the genus Saccamoeba is paraphyletic and is still in need of revision as it currently occupies two distinct lineages. Here, we report a new clade on the tree of Tubulinea, which we infer represents a novel genus that we name Ptolemeba n. gen. This genus subsumes a clade of hartmannellid amoebae that were previously considered in the genus Saccamoeba, but whose mitochondrial morphology is distinct from Saccamoeba. In accordance with previous research, we formalize the clade as distinct from Saccamoeba. Transmission electron microscopy of our isolates illustrate that both molecularly discrete species can be further differentiated by their unique mitochondrial cristal morphology.     

A standardized reanalysis of molecular phylogenetic hypotheses of Gobioidei

Gobioidei is a suborder of perciform fishes with about 2000 species distributed worldwide. Despite the evolutionary and ecological importance of gobioids, their phylogenetic inter- and intrarelationships are still poorly understood. Only a few studies (either morphological or molecular) have tackled the phylogeny of Gobioidei as a whole. Of these, only six studies thus far have addressed gobioid intrarelationships based on molecular data (each using different taxon sampling, genes, outgroups and method of phylogenetic inference), yielding contrasting results regarding the phylogenetic relationships among major lineages. In this study, we have reanalysed data from four of these molecular phylogenetic studies of Gobioidei under standardization criteria (same outgroup and methods of phylogenetic inference) in order to assess the robustness of their results, as well as to identify which parts of the gobioid tree are least resolved. Results from all datasets reanalysed in this study are generally similar to those of the respective original studies, and suggest broad patterns of phylogenetic relationships among gobioid lineages. However, there are numerous topological discrepancies among the four studies, support is low for many phylogenetic relationships and topology tests are unable to reject the vast majority of alternative topologies tested. The concatenation of datasets yields a relatively robust phylogeny of major lineages of Gobioidei, but there are some issues of overlap and missing data, which are ameliorated with the inclusion of additional homologous sequences from GenBank that increase dataset completeness. Because both monophyly of major gobioid groups and phylogenetic relationships among them cannot be fully resolved, it is clear that further phylogenetic research is needed, and this should be accompanied by a major taxonomic revision of the Gobioidei. Nevertheless, even with the relatively unstable nature of the available molecular phylogenies, there are some monophyletic units that can be identified, and a basic structure of the gobioid tree appears evident.     

Phylogeny of New Zealand hepialid moths (Lepidoptera: Hepialidae) inferred from a cladistic analysis of morphological data

The phylogeny of the New Zealand hepialid moths was estimated from a cladistic analysis of sixty‐three morphological characters, from all life cycle stages. One hundred and sixteen maximum parsimony trees were produced. The phylogenetic reconstruction indicated that the currently recognized generic concepts, and the four informal lineages hypothesized in a previous morphological taxonomic revision, were monophyletic. The relationships of species within genus Wiseana were not fully resolved. Analysis of a data set of thirty‐nine adult male characters from the New Zealand taxa and the Australian genera Jeana, Oxycanus and Trictena supported the monophyly of the New Zealand ‘Oxycanus’ s.s lineage.     

Phylogenetic analyses reveal reliable morphological markers to classify mega‐diversity in Onthophagini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae)

Based largely on homoplastic characters of external morphology, the current systematics of the tribe Onthophagini and allied dung beetle lineages is unstable, contradictory, and thus inefficient. A number of recently proposed molecular phylogenies conflict strongly with each other and with formal classification, and none of them provides new tools for the improvement of dung beetle systematics. We explored the source of these inconsistencies by performing an independent, morphology‐based phylogenetic analysis of the “Serrophorus complex”, one of the most systematically confusing knots among the onthophagines, that involves 52 species from various genera of Onthophagini and allied tribes. The phylogenetic pattern revealed conflicts with existing classifications and with most of the earlier molecular phylogenies. However, it was largely congruent with the molecular phylogeny (Evolution 2005, 59 , 1060), using the largest gene sampling thus far. All current competing phylogenetic hypotheses were evaluated against each other, and the degree of their biogeographic plausibility was used as an additional evaluative criterion. Of the 91 morphological characters involved in our analyses, traits belonging to the endophallic sclerites of the aedeagus had a very strong phylogenetic signal. Terminology of these endophallic characters was established and their morphology was studied in detail, illustrated, and presented as a tool for further practical use. The enormous variety of shapes of the lamella copulatrix within the Onthophagini and allies present a methodological problem in character coding for phylogenetic analyses. Based on the performance of alternative coding approaches, it is argued that a seemingly less informative absence/presence coding scheme would be a better choice. The phylogenetic structure of the Serrophorus complex has been largely resolved, and some taxonomic changes improving its systematics are recommended.
© The Willi Hennig Society 2011.     

Navigating the mtDNA road map out of the morphological maze: interpreting morphological variation in the diverse Monomorium rothsteini (Forel) complex (Hymenoptera: Formicidae)

Monomorium is a large and diverse ant genus with speciose radiations in both the Afrotropical and Australian regions. According to the most recent taxonomic revision, many Australian species are characterised by very broad distributions and variable morphology, which suggests that some species may be unrecognised species complexes. With a continent‐wide distribution and diverse yet overlapping morphology, M. rothsteini (Forel) is representative of the greater challenge that exists in Australian Monomorium systematics. Here we investigate species boundaries in M. rothsteini using a molecular phylogenetic framework to interpret the complex overlap of nine morphological characters (with 31 states) and examine biogeographic relationships among the lineages. Bayesian inference resolved 38 mtDNA lineages that were morphologically separable, at least from their sister lineage. Although the morphological characters were intermixed across the phylogeny, instances of inseparable morphology among sister clades was rare. Seventeen lineages exhibited complete morphological overlap with one or more other lineages and could not be separated by Principal Component Analysis based on 12 morphometric variables. Two‐thirds of all lineages occurred sympatrically with one or more both genetically and morphologically divergent lineages. The two nuclear markers EF1αF2 and wingless were used to generate haplotype networks which were characterised by a star‐like pattern indicative of a rapid and recent radiation. Several haplotypes for both nuclear gene regions were shared among individuals occurring in separate mtDNA clades which we were also unable to distinguish morphologically or that were occurring in sympatry, indicating possible introgression in both the mtDNA and nuclear genomes. Clear biogeographic affinities among samples within a lineage were detected but there was no overall pattern in the biogeographic relationships among the lineages. We conclude that M. rothsteini is a large species complex that has undergone a complex evolutionary history following aridification of the Australian continent, and discuss the implications of this conclusion for the systematics of Australian Monomorium more generally.     

A single origin for the limnetic–euryhaline taxa in the Corbulidae (Bivalvia)

The bivalve family Corbulidae, known colloquially as ‘basket clams’, includes species tolerating a wide variety of habitats ranging from open marine to freshwater. Previous studies of corbulid phylogenetics have been based mainly on shell morphology and to some extent soft tissue anatomy. However, these studies have been inadequate for corbulid classification because of difficulties in determining the inter‐relationships of primarily marine species with non‐marine species, the latter commonly exhibiting highly divergent morphological, ecological and environmental characteristics from their marine counterparts. The first molecular phylogenetic study of the Corbulidae is presented herein, analysing DNA sequences from the 18S rRNA and 28S rRNA genes, separately and in combination. Fifteen corbulid species and 14 outgroup taxa were included in the analyses. Corbulidae is resolved as monophyletic, comprising three groups with varying support. The non‐marine species form one group that we name as the subclade ‘limnetic–euryhaline Corbulidae’ (LEC) and comprising the genera Lentidium, Erodona and Potamocorbula. This LEC, which is consistently recovered as monophyletic, is globally distributed. The marine Corbulidae are divided into two well‐supported lineages in combined analyses although there are inconsistencies in their membership between single‐gene analyses. One of the two lineages consists of primarily Western Pacific taxa and the other of North American and Caribbean taxa. Finally, the authors advocate further study on the LEC to mitigate potential biological invasions beyond their native distribution.     

Assessing spatial and environmental drivers of phylogenetic structure in Brazilian Araucaria forests

Evidence of phylogenetic conservatism in plant ecological traits has accumulated over the past few years, suggesting an interplay between the distribution of phylogenetic clades and major environmental gradients. Nonetheless, determining what environmental factors underlie the distribution of phylogenetic lineages remains a challenge because environmental factors are correlated with spatial gradients where the latter might indicate some degree of dispersal limitation in phylogenetic pools. We analyzed the phylogenetic structure of plant assemblages across the Brazilian Araucaria forests and assessed how phylogenetic structure responds to environmental and spatial gradients. We compiled data on plant occurrence in 45 plots across the Araucaria forest biome. The phylogenetic structure of the plots was characterized using phylogenetic fuzzy‐weighting followed by principal coordinates of phylogenetic structure (PCPS). We used distance‐based redundancy analysis (db‐RDA) to analyze the relationships between phylogenetic clades and environmental and spatial factors. Variation partitioning showed that the phylogenetic structure of Brazilian Araucaria forests was better explained by environment factors (altitude and annual mean temperature) than by space. Yet, spatially‐structured environmental variation explained about one‐third of total variation in the phylogenetic structure. Thus, the influence of spatial filters on the phylogenetic structure was more related to environmental gradients across the Brazilian Araucaria forest biome than to dispersal limitation of phylogenetic lineages. Furthermore, the influence of explanatory factors on the phylogenetic structure was concentrated in few nodes, the one splitting tree ferns from seed plants, and a second splitting malvids from other eurosids. Assessing the functional links between species distribution patterns and environmental gradients is not an easy task when we have to deal with large species pools. Identifying major phylogenetic gradients across an environmental and/or geographical range of interest can represent a first step towards a better understanding of general assembly processes in ecological communities.     

Phylogenetic and biogeographic complexity of Magnoliaceae in the Northern Hemisphere inferred from three nuclear data sets

This study employs three nuclear genes (PHYA, LFY, and GAI1) to reconstruct the phylogenetic and biogeographic history of Magnoliaceae. A total of 104 samples representing 86 taxa from all sections and most subsections were sequenced. Twelve major groups are well supported to be monophyletic within Magnoliaceae and these groups are largely consistent with the recent taxonomic revision at the sectional and subsectional levels. However, relationships at deeper nodes of the subfamily Magnolioideae remain not well resolved. A relaxed clock relying on uncorrelated rates suggests that the complicated divergent evolution of Magnolioideae began around the early Eocene (54.57mya), concordant with paleoclimatic and fossil evidence. Intercontinental disjunctions of Magnoliaceae in the Northern Hemisphere appear to have originated during at least two geologic periods. Some occurred after the middle Miocene, represented by two well-recognized temperate lineages disjunct between eastern Asia and eastern North America. The others may have occurred no later than the Oligocene, with ancient separations between or within tropical and temperate lineages.     

Probing the Monophyly of the Sphaeropleales (Chlorophyceae) Using Data From Five Genes

Molecular phylogenetic analyses have had a major impact on the classification of the green algal class Chlorophyceae, corroborating some previous evolutionary hypotheses, but primarily promoting new interpretations of morphological evolution. One set of morphological traits that feature prominently in green algal systematics is the absolute orientation of the flagellar apparatus in motile cells, which correlates strongly with taxonomic classes and orders. The order Sphaeropleales includes diverse green algae sharing the directly opposite (DO) flagellar apparatus orientation of their biflagellate motile cells. However, algae across sphaeroplealean families differ in specific components of the DO flagellar apparatus, and molecular phylogenetic studies often have failed to provide strong support for the monophyly of the order. To test the monophyly of Sphaeropleales and of taxa with the DO flagellar apparatus, we conducted a molecular phylogenetic study of 16 accessions representing all known families and diverse affiliated lineages within the order, with data from four plastid genes (psaA, psaB, psbC, rbcL) and one nuclear ribosomal gene (18S). Although single‐gene analyses varied in topology and support values, analysis of combined data strongly supported a monophyletic Sphaeropleales. Our results also corroborated previous phylogenetic hypotheses that were based on chloroplast genome data from relatively few taxa. Specifically, our data resolved Volvocales, algae possessing predominantly biflagellate motile cells with clockwise (CW) flagellar orientation, as the monophyletic sister lineage to Sphaeropleales, and an alliance of Chaetopeltidales, Chaetophorales, and Oedogoniales, orders having multiflagellate motile cells with distinct flagellar orientations involving the DO and CW forms.     

Phylogenetics, biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): effects of character exclusion, data partitioning and missing data

The phylogenetic relationships, biogeography and classification of, and morpho‐behavioral (M/B) evolution in, gamebirds (Aves: Galliformes) are investigated. In‐group taxa (rooted on representatives of the Anseriformes) include 158 species representing all suprageneric galliform taxa and 65 genera. The characters include 102 M/B attributes and 4452 nucleic acid base pairs from mitochondrial cytochrome b (CYT B), NADH dehydrogenase subunit 2 (ND2), 12S ribosomal DNA (12S) and control region (CR), and nuclear ovomucoid intron G (OVO‐G). Analysis of the combined character data set yielded a single, completely resolved cladogram that had the highest levels of jackknife support, which suggests a need for a revised classification for the phasianine galliforms. Adding 102 M/B characters to the combined CYT B and ND2 partitions (2184 characters) decisively overturns the topology suggested by analysis of the two mtDNA partitions alone, refuting the view that M/B characters should be excluded from phylogenetic analyses because of their relatively small number and putative character state ambiguity. Exclusion of the OVO‐G partition (with > 70% missing data) from the combined data set had no effect on cladistic structure, but slightly lowered jackknife support at several nodes. Exclusion of third positions of codons in an analysis of a CYT B + ND2 partition resulted in a massive loss of resolution and support, and even failed to recover the monophyly of the Galliformes with jackknife support. A combined analysis of putatively less informative, “non‐coding” characters (CYT B/ND2 third position sites + CR +12S + OVO‐G sequences) yielded a highly resolved consensus cladogram congruent with the combined‐evidence cladogram. Traditionally recognized suprageneric galliform taxa emerging in the combined cladogram are: the families Megapodiidae (megapodes), Cracidae (cracids), Numididae (guineafowls), Odontophoridae (New World quails) and Phasianidae (pheasants, pavonines, partridges, quails, francolins, spurfowls and grouse) and the subfamilies Cracinae (curassows, chachalacas and the horned guan), Penelopinae (remaining guans), Pavoninae sensu lato (peafowls, peacock pheasants and argus pheasants), Tetraoninae (grouse) and Phasianinae (pheasants minus Gallus). The monophyly of some traditional groupings (e.g., the perdicinae: partridges/quails/francolins) is rejected decisively, contrasted by the emergence of other unexpected groupings. The most remarkable phylogenetic results are the placement of endemic African galliforms as sisters to geographically far‐distant taxa in Asia and the Americas. Biogeographically, the combined‐data cladogram supports the hypothesis that basal lineages of galliforms diverged prior to the Cretaceous/Tertiary (K‐T) Event and that the subsequent cladogenesis was influenced by the break‐up of Gondwana. The evolution of gamebirds in Africa, Asia and the Americas has a far more complicated historical biogeography than suggested to date. With regard to character evolution: spurs appear to have evolved at least twice within the Galliformes; a relatively large number of tail feathers (≥ 14) at least three times; polygyny at least twice; and sexual dimorphism many times. © The Willi Hennig Society 2006.     

Molecular phylogeny and character mapping support generic adjustments in the Tetrapteroid clade (Malpighiaceae)

In the past two decades, Malpighiaceae has gone through unprecedented changes in classification due to molecular phylogenetic studies. However, some generic adjustments are still needed to make all genera monophyletic. In this study, we propose the new genus Glicophyllum to accommodate the species of Tetrapterys pro parte, a well‐supported lineage recovered by us and by the most recent phylogenetic studies of the family. Additionally, we propose the synonymy of Aenigmatanthera under Niedenzuella to finally render the latter monophyletic. Morphological characters related to inflorescence architecture, bract and bracteole size, presence of glands on the bracts and bracteoles, style size and thickness, and ratio length among superior and anterior lateral wings of mericarps are important for circumscribing these lineages.     

MOLECULAR PHYLOGENY OF SELECTED SPECIES OF THE ORDER DINOPHYSIALES (DINOPHYCEAE)—TESTING THE HYPOTHESIS OF A DINOPHYSIOID RADIATION1

Almost 80 years ago, a radiation scheme based on structural resemblance was first outlined for the marine order Dinophysiales. This hypothetical radiation illustrated the relationship between the dinophysioid genera and included several independent, extant lineages. Subsequent studies have supplied additional information on morphology and ecology to these evolutionary lineages. We have for the first time combined morphological information with molecular phylogenies to test the dinophysioid radiation hypothesis in a modern context. Nuclear‐encoded LSU rDNA sequences including domains D1‐D6 from 27 species belonging to Dinophysis Ehrenb., Ornithocercus F. Stein, Phalacroma F. Stein, Amphisolenia F. Stein, Citharistes F. Stein, and Histioneis F. Stein were obtained from the Indian Ocean. Previously, LSU rDNA has only been determined from one of these. In Bayesian analyses, Amphisolenia formed a long basal clade to the other dinophysioids. These diverged into two separate lineages, the first comprised species with a classical Phalacroma outline, also including the type species P. porodictyum F. Stein. Thus, we propose to reinstate the genus Phalacroma. The relationship between the genera in the second lineage was not well resolved. However, the molecular phylogeny supported monophyly of Histioneis and Citharistes and showed the genus Dinophysis to be polyphyletic and in need of a taxonomic revision. Species of Ornithocercus grouped with Citharistes, but this relationship remained unresolved. The phylogenetic trees furthermore revealed convergent evolution of several morphological characters in the dinophysioids. According to the molecular data, the dinophysioids appeared to have evolved quite differently from the radiation schemes previously hypothesized. Four dinophysioid species had identical LSU rDNA sequences to other well‐established species.     

Phylogeny of Coreopsideae (Asteraceae) using ITS sequences suggests lability in reproductive characters

Relationships among the 21 genera within the tribe Coreopsideae (Asteraceae) remain poorly resolved despite phylogenetic studies using morphological and anatomical traits. Recent molecular phylogenies have also indicated that some Coreopsideae genera are not monophyletic. We used internal transcribed spacer (ITS) sequences from representatives of 19 genera, as well as all major lineages in those genera that are not monophyletic, to examine phylogenetic relationships within this group. To examine the affects of alignment and method of analysis on our conclusions, we obtained alignments using five different parameters and analyzed all five alignments with distance, parsimony, and Bayesian methods. The method of analysis had a larger impact on relationships than did alignments, although different analytical methods gave very similar results. Although not all relationships could be resolved, a number of well-supported lineages were found, some in conflict with earlier hypotheses. We did not find monophyly in Bidens, Coreopsis, and Coreocarpus, though other genera were monophyletic for the taxa we included. Morphological and anatomical traits which have been used previously to resolve phylogenetic relationships in this group were mapped onto the well-supported nodes of the ITS phylogeny. This analysis indicated that floral and fruit characters, which have been used extensively in phylogenetic studies in the Coreopsideae, show a higher degree of evolutionary lability in this group than the more highly conserved vegetative and photosynthetic traits.     

Systematic revision of mountain dragons (Reptilia: Agamidae: Diploderma) in China,with descriptions of six new species and discussion on their conservation

Cryptic diversity not only introduces confusion to taxonomic studies, but it also poses major challenges to conservation and environmental legislation. One such troubling group are the Mountain Dragons in the genus Diploderma in Southwest China. Previous studies have suggested that the genus contains considerable cryptic diversity, particularly in the D. flaviceps complex. Owing to taxonomic confusion, micro‐endemic lineages are still neglected by the Chinese wildlife protection laws, despite their urgent conservation needs. Combining multivariate morphological and multi‐locus phylogenetic data, we provide the first integrative systematic revision of the genus Diploderma. Specifically, we confirm that the six examined populations of D. cf. flaviceps from the upper Jinsha and Yalong River Valleys in Sichuan and Yunnan Provinces represent six cryptic, undescribed lineages, and we describe each of them as a new species. With the updated taxonomy and distribution information, we discuss the taxonomy of the D. flaviceps complex in Southwest China, provide an updated diagnostic key along with distributional ranges for all species of the genus, and discuss some of the suspicious records of other congeners in China. Lastly, we evaluate the IUCN status of each of the six new species and highlight the major challenges for Diploderma conservation in China due to delayed environmental legislation and misleading conservation assessments.     

Phylogenetic relationships of Microdontinae (Diptera: Syrphidae) based on molecular and morphological characters

The intrasubfamilial classification of Microdontinae Rondani (Diptera: Syrphidae) has been a challenge: until recently more than 300 out of more than 400 valid species names were classified in Microdon Meigen. We present phylogenetic analyses of molecular and morphological characters (both separate and combined) of Microdontinae. The morphological dataset contains 174 characters, scored for 189 taxa (9 outgroup), representing all 43 presently recognized genera and several subgenera and species groups. The molecular dataset, representing 90 ingroup species of 28 genera, comprises sequences of five partitions in total from the mitochondrial gene COI and the nuclear ribosomal genes 18S and 28S. We test the sister‐group relationship of Spheginobaccha with the other Microdontinae, attempt to elucidate phylogenetic relationships within the Microdontinae and discuss uncertainties in the classification of Microdontinae. Trees based on molecular characters alone are poorly resolved, but combined data are better resolved. Support for many deeper nodes is low, and placement of such nodes differs between parsimony and Bayesian analyses. However, Spheginobaccha is recovered as highly supported sister group in both. Both analyses agree on the early branching of Mixogaster, Schizoceratomyia, Afromicrodon and Paramicrodon. The taxonomical rank in relation to the other Syrphidae is discussed briefly. An additional analysis based on morphological characters only, including all 189 taxa, used implied weighting. A range of weighting strengths (k‐values) is applied, chosen such that values of character fit of the resulting trees are divided into regular intervals. Results of this analysis are used for discussing the phylogenetic relationships of genera unrepresented in the molecular dataset.     

Remarkable ancient divergences amongst neglected lorisiform primates

Lorisiform primates (Primates: Strepsirrhini: Lorisiformes) represent almost 10% of the living primate species and are widely distributed in sub‐Saharan Africa and South/South‐East Asia; however, their taxonomy, evolutionary history, and biogeography are still poorly understood. In this study we report the largest molecular phylogeny in terms of the number of represented taxa. We sequenced the complete mitochondrial cytochrome b gene for 86 lorisiform specimens, including ～80% of all the species currently recognized. Our results support the monophyly of the Galagidae, but a common ancestry of the Lorisinae and Perodicticinae (family Lorisidae) was not recovered. These three lineages have early origins, with the Galagidae and the Lorisinae diverging in the Oligocene at about 30 Mya and the Perodicticinae emerging in the early Miocene. Our mitochondrial phylogeny agrees with recent studies based on nuclear data, and supports Euoticus as the oldest galagid lineage and the polyphyletic status of Galagoides. Moreover, we have elucidated phylogenetic relationships for several species never included before in a molecular phylogeny. The results obtained in this study suggest that lorisiform diversity remains substantially underestimated and that previously unnoticed cryptic diversity might be present within many lineages, thus urgently requiring a comprehensive taxonomic revision of this primate group. © 2015 The Linnean Society of London